Greenhouse gas emissions from African lakes are no longer a blind spot.

Natural lakes are thought to be globally important sources of greenhouse gases (CO2, CH4, and N2O) to the atmosphere although nearly no data have been previously reported from Africa. We collected CO2, CH4, and N2O data in 24 African lakes that accounted for 49% of total lacustrine surface area of the African continent and covered a wide range of morphology and productivity. The surface water concentrations of dissolved CO2 were much lower than values attributed in current literature to tropical lakes and lower than in boreal systems because of a higher productivity. In contrast, surface water-dissolved CH4 concentrations were generally higher than in boreal systems. The lowest CO2 and the highest CH4 concentrations were observed in the more shallow and productive lakes. Emissions of CO2 may likely have been substantially overestimated by a factor between 9 and 18 in African lakes and between 6 and 26 in pan-tropical lakes.

Microbial eukaryote assemblages and potential novel diversity in four tropical East African Great Lakes.

East African Great Lakes are old and unique natural resources heavily utilized by their bordering countries. In those lakes, ecosystem functioning is dominated by pelagic processes, where microorganisms are key components; however, protistan diversity is barely known. We investigated the community composition of small eukaryotes (<10 µm) in surface waters of four African Lakes (Kivu, Edward, Albert and Victoria) by sequencing the 18S rRNA gene. Moreover, in the meromictic Lake Kivu, two stations were vertically studied. We found high protistan diversity distributed in 779 operational taxonomic units (OTUs), spanning in 11 high-rank lineages, being Alveolata (31%), Opisthokonta (20%) and Stramenopiles (17%) the most represented supergroups. Surface protistan assemblages were associated with conductivity and productivity gradients, whereas depth had a strong effect on protistan community in Kivu, with higher contribution of heterotrophic organisms. Approximately 40% of OTUs had low similarity (<90%) with reported sequences in public databases; these were mostly coming from deep anoxic waters of Kivu, suggesting a high extent of novel diversity. We also detected several taxa so far considered exclusive of marine ecosystems. Our results unveiled a complex and largely undescribed protistan community, in which several lineages have adapted to different niches after crossing the salinity boundary.

First Report on Microcystis as a Potential Microviridin Producer in Bulgarian Waterbodies.

Bulgaria, situated on the Balkan Peninsula, is rich in small and shallow, natural and man-made non-lotic waterbodies, which are threatened by blooms of Cyanoprokaryota/Cyanobacteria. Although cyanotoxins in Bulgarian surface waters are receiving increased attention, there is no information on microviridins and their producers. This paper presents results from a phytoplankton study, conducted in August 2019 in three lakes (Durankulak, Vaya, Uzungeren) and five reservoirs (Duvanli, Mandra, Poroy, Sinyata Reka, Zhrebchevo) in which a molecular-genetic analysis (PCR based on the precursor mdnA gene and subsequent translation to amino acid alignments), combined with conventional light microscopy and an HPLC analysis of marker pigments, were applied for the identification of potential microviridin producers. The results provide evidence that ten strains of the genus Microcystis, and of its most widespread species M. aeruginosa in particular, are potentially toxigenic in respect to microviridins. The mdnA sequences were obtained from all studied waterbodies and their translation to amino-acid alignments revealed the presence of five microviridin variants (types B/C, Izancya, CBJ55500.1 (Microcystis 199), and MC19, as well as a variant, which was very close to type A). This study adds to the general understanding of the microviridin occurrence, producers, and sequence diversity.

The possible occurrence of iron-dependent anaerobic methane oxidation in an Archean Ocean analogue.

In the ferruginous and anoxic early Earth oceans, photoferrotrophy drove most of the biological production before the advent of oxygenic photosynthesis, but its association with ferric iron (Fe3+) dependent anaerobic methane (CH4) oxidation (AOM) has been poorly investigated. We studied AOM in Kabuno Bay, a modern analogue to the Archean Ocean (anoxic bottom waters and dissolved Fe concentrations > 600 µmol L-1). Aerobic and anaerobic CH4 oxidation rates up to 0.12 ± 0.03 and 51 ± 1 µmol L-1 d-1, respectively, were put in evidence. In the Fe oxidation-reduction zone, we observed high concentration of Bacteriochlorophyll e (biomarker of the anoxygenic photoautotrophs), which co-occurred with the maximum CH4 oxidation peaks, and a high abundance of Candidatus Methanoperedens, which can couple AOM to Fe3+ reduction. In addition, comparison of measured CH4 oxidation rates with electron acceptor fluxes suggest that AOM could mainly rely on Fe3+ produced by photoferrotrophs. Further experiments specifically targeted to investigate the interactions between photoferrotrophs and AOM would be of considerable interest. Indeed, ferric Fe3+-driven AOM has been poorly envisaged as a possible metabolic process in the Archean ocean, but this can potentially change the conceptualization and modelling of metabolic and geochemical processes controlling climate conditions in the Early Earth.

Long-term impacts of nutrient control, climate change, and invasive clams on phytoplankton and cyanobacteria biomass in a large temperate river.

Recent studies suggest that climate change, with warmer water temperatures and lower and longer low flows, may enhance harmful planktic cyanobacterial growth in lakes and large rivers. Concomitantly, controlling nutrient loadings has proven effective in reducing phytoplankton biomass especially in North America and Western Europe. In addition, the impact of invasive benthic filter-feeder species such as Corbicula on phytoplankton has largely been overlooked in large rivers, leading to even more uncertainty in predicting future trajectories in river water quality. To investigate how nutrient control, climate change and invasion of benthic filter-feeders may affect phytoplankton biomass and composition, we assembled a large database on the entire water course of the River Loire (France) over three decades (1991-2019). We focus on cyanobacteria to provide an in-depth analysis of the 30-year trend and insights on future possible trajectories. Since 1991, total phytoplankton and cyanobacteria biomasses have decreased 10-fold despite warmer water temperature (+0.23 °C·decade-1) and lower summer flow (-0.25 L·s-1·km-2·decade-1). In the long-term, the contribution of planktic cyanobacteria to total biomass was on average 2.8%. The main factors driving total phytoplankton and cyanobacteria biomasses were total phosphorus (4-fold decrease), the abundance of Corbicula clams (from absence before 1998 to 250-1250 individuals·m-2 after 2010), the duration of summer low flows and the intensity of summer heatwaves. The River Loire constitutes an example in Europe of how nutrient control can be an efficient mitigation strategy, counteracting already visible effects of climate change on the thermal regime and flow pattern of the river. This may hold true under future conditions, but further work is needed to account for the climate trajectory, land and water use scenarios, the risk of enhanced benthic biofilm and macrophyte proliferation, together with the spread of invasive filter-feeding bivalves.

Long-term trends in trait structure of riverine communities facing predation risk increase and trophic resource decline.

Many large European rivers have undergone multiple pressures that have strongly impaired ecosystem functioning at different spatial and temporal scales. Global warming and other environmental changes have favored the success of invasive species, deeply modifying the structure of aquatic communities in large rivers. Some exogenous species could alter trophic interactions within assemblages by increasing the predation risk for potential prey species (top-down effect) and limiting the dynamics of others via resource availability limitation (bottom-up effect). Furthermore, large transboundary rivers are complex aquatic ecosystems that have often been poorly investigated so that data for assessing long-term ecological trends are missing. In this study, we propose an original approach for investigating long-term combined effects of global warming, trophic resource decrease, predation risk, and water quality variations on the trait-based structure of macroinvertebrate and fish assemblages over 26 yr (1985-2011) and 427-km stretch of the river Meuse (France and Belgium). The study of temporal variations in biological, physiological, and ecological traits of macroinvertebrate and fish allowed identifying community trends and distinguishing impacts of environmental perturbations from those induced by biological alterations. We provide evidence, for this large European river, of an increase in water temperature (close to 1°C) and a decrease in phytoplankton biomass (-85%), as well as independent effects of these changes on both invertebrate and fish communities. The reduction of trophic resources in the water column by invasive molluscs has dramatically affected the density of omnivorous fish in favor of invertebrate feeders, while scrapers became the major feeding guild among invertebrates. Macroinvertebrate and fish communities have shifted from large-sized organisms with low fecundity to prolific, small-sized organisms, with early maturity, as a response to increased predation pressure.

Pelagic photoferrotrophy and iron cycling in a modern ferruginous basin.

Iron-rich (ferruginous) ocean chemistry prevailed throughout most of Earth's early history. Before the evolution and proliferation of oxygenic photosynthesis, biological production in the ferruginous oceans was likely driven by photoferrotrophic bacteria that oxidize ferrous iron {Fe(II)} to harness energy from sunlight, and fix inorganic carbon into biomass. Photoferrotrophs may thus have fuelled Earth's early biosphere providing energy to drive microbial growth and evolution over billions of years. Yet, photoferrotrophic activity has remained largely elusive on the modern Earth, leaving models for early biological production untested and imperative ecological context for the evolution of life missing. Here, we show that an active community of pelagic photoferrotrophs comprises up to 30% of the total microbial community in illuminated ferruginous waters of Kabuno Bay (KB), East Africa (DR Congo). These photoferrotrophs produce oxidized iron {Fe(III)} and biomass, and support a diverse pelagic microbial community including heterotrophic Fe(III)-reducers, sulfate reducers, fermenters and methanogens. At modest light levels, rates of photoferrotrophy in KB exceed those predicted for early Earth primary production, and are sufficient to generate Earth's largest sedimentary iron ore deposits. Fe cycling, however, is efficient, and complex microbial community interactions likely regulate Fe(III) and organic matter export from the photic zone.

Vertical Distribution of Functional Potential and Active Microbial Communities in Meromictic Lake Kivu.

The microbial community composition in meromictic Lake Kivu, with one of the largest CH4 reservoirs, was studied using 16S rDNA and ribosomal RNA (rRNA) pyrosequencing during the dry and rainy seasons. Highly abundant taxa were shared in a high percentage between bulk (DNA-based) and active (RNA-based) bacterial communities, whereas a high proportion of rare species was detected only in either an active or bulk community, indicating the existence of a potentially active rare biosphere and the possible underestimation of diversity detected when using only one nucleic acid pool. Most taxa identified as generalists were abundant, and those identified as specialists were more likely to be rare in the bulk community. The overall number of environmental parameters that could explain the variation was higher for abundant taxa in comparison to rare taxa. Clustering analysis based on operational taxonomic units (OTUs at 0.03 cutoff) level revealed significant and systematic microbial community composition shifts with depth. In the oxic zone, Actinobacteria were found highly dominant in the bulk community but not in the metabolically active community. In the oxic-anoxic transition zone, highly abundant potentially active Nitrospira and Methylococcales were observed. The co-occurrence of potentially active sulfur-oxidizing and sulfate-reducing bacteria in the anoxic zone may suggest the presence of an active yet cryptic sulfur cycle.

Bacterial community composition in three freshwater reservoirs of different alkalinity and trophic status.

In order to investigate the factors controlling the bacterial community composition (BCC) in reservoirs, we sampled three freshwater reservoirs with contrasted physical and chemical characteristics and trophic status. The BCC was analysed by 16S rRNA gene amplicon 454 pyrosequencing. In parallel, a complete dataset of environmental parameters and phytoplankton community composition was also collected. BCC in the analysed reservoirs resembled that of epilimnetic waters of natural freshwater lakes with presence of Actinobacteria, Alpha- and Betaproteobacteria, Cytophaga-Flavobacteria-Bacteroidetes (CFB) and Verrucomicrobia groups. Our results evidenced that the retrieved BCC in the analysed reservoirs was strongly influenced by pH, alkalinity and organic carbon content, whereas comparatively little change was observed among layers in stratified conditions.

Carbon cycling of Lake Kivu (East Africa): net autotrophy in the epilimnion and emission of CO2 to the atmosphere sustained by geogenic inputs.

We report organic and inorganic carbon distributions and fluxes in a large (>2000 km2) oligotrophic, tropical lake (Lake Kivu, East Africa), acquired during four field surveys, that captured the seasonal variations (March 2007-mid rainy season, September 2007-late dry season, June 2008-early dry season, and April 2009-late rainy season). The partial pressure of CO2 (pCO2) in surface waters of the main basin of Lake Kivu showed modest spatial (coefficient of variation between 3% and 6%), and seasonal variations with an amplitude of 163 ppm (between 579±23 ppm on average in March 2007 and 742±28 ppm on average in September 2007). The most prominent spatial feature of the pCO2 distribution was the very high pCO2 values in Kabuno Bay (a small sub-basin with little connection to the main lake) ranging between 11,213 ppm and 14,213 ppm (between 18 and 26 times higher than in the main basin). Surface waters of the main basin of Lake Kivu were a net source of CO2 to the atmosphere at an average rate of 10.8 mmol m(-2) d(-1), which is lower than the global average reported for freshwater, saline, and volcanic lakes. In Kabuno Bay, the CO2 emission to the atmosphere was on average 500.7 mmol m(-2) d(-1) (∼46 times higher than in the main basin). Based on whole-lake mass balance of dissolved inorganic carbon (DIC) bulk concentrations and of its stable carbon isotope composition, we show that the epilimnion of Lake Kivu was net autotrophic. This is due to the modest river inputs of organic carbon owing to the small ratio of catchment area to lake surface area (2.15). The carbon budget implies that the CO2 emission to the atmosphere must be sustained by DIC inputs of geogenic origin from deep geothermal springs.

Decadal trends and common dynamics of the bio-optical and thermal characteristics of the African Great Lakes.

The Great Lakes of East Africa are among the world's most important freshwater ecosystems. Despite their importance in providing vital resources and ecosystem services, the impact of regional and global environmental drivers on this lacustrine system remains only partially understood. We make a systematic comparison of the dynamics of the bio-optical and thermal properties of thirteen of the largest African lakes between 2002 and 2011. Lake surface temperatures had a positive trend in all Great Lakes outside the latitude of 0° to 8° south, while the dynamics of those lakes within this latitude range were highly sensitive to global inter-annual climate drivers (i.e. El Niño Southern Oscillation). Lake surface temperature dynamics in nearly all lakes were found to be sensitive to the latitudinal position of the Inter Tropical Convergence Zone. Phytoplankton dynamics varied considerably between lakes, with increasing and decreasing trends. Intra-lake differences in both surface temperature and phytoplankton dynamics occurred for many of the larger lakes. This inter-comparison of bio-optical and thermal dynamics provides new insights into the response of these ecosystems to global and regional drivers.

Phylogeny and androgenesis in the invasive Corbicula clams (Bivalvia, Corbiculidae) in Western Europe.

BACKGROUND: The genus Corbicula is one of the most invasive groups of molluscs. It includes both sexual and androgenetic lineages. The present study re-assessed the different morphotypes and haplotypes of West European Corbicula in order to clarify their taxonomic identification and phylogenetic relationships with American and Asian Corbicula clams. We studied several populations from West European river basins (Meuse, Seine, Rhine and Rhône) through an "integrative taxonomy" approach. We combined morphology, partial mitochondrial COI and cyt b sequences and eleven microsatellite loci. Furthermore, we looked for discrepancies between mtDNA and nrDNA/morphology, indicative of androgenesis between lineages. RESULTS: There are three Corbicula morphotypes in Western Europe associated to three mitochondrial lineages and three genotypes. Form R shares the same COI haplotype as the American form A and the Japanese C. leana. Form S and the American form C have the same haplotype, although their morphologies seem divergent. The European form Rlc belongs to the same mitochondrial lineage as both the American form B and the Asian C. fluminea.Interestingly, within each haplotype/genotype or lineage, no genetic diversity was found although their invasive success is high. Moreover, we detected rare mismatches between mtDNA and nrDNA/morphology, indicative of androgenesis and mitochondrial capture between form R and form S and therefore challenging the phylogenetic relatedness and the species status within this genus. The global phylogenetic analysis revealed that the sexual Corbicula lineages seem restricted to the native areas while their androgenetic relatives are widespread and highly invasive. CONCLUSIONS: We clarified the discrepancies and incongruent results found in the literature about the European morphotypes of Corbicula and associated mitochondrial lineages. The three West European morphotypes belong to three distinct nuclear and mitochondrial lineages. However mitochondrial capture occurs in sympatric populations of forms R and S. The species status of the morphotypes therefore remains doubtful. Moreover the androgenetic lineages seem widely distributed compared to their sexual relatives, suggesting that androgenesis and invasive success may be linked in the genus Corbicula.

Protist herbivory: a key pathway in the pelagic food web of Lake Tanganyika.

Herbivory and bacterivory by phagotrophic protists were estimated in the southern basin of the oligotrophic Lake Tanganyika at different seasons (in the rainy season in February-March 2007 and in the dry season in July-August 2006 and September 2007), using two independent methods: the selective inhibitor technique for assessing community grazing on picocyanobacteria (PCya) and fluorescently labelled bacteria (FLB) and Synechococcus (FLA) to estimate bacterivory and herbivory by phagotrophic nanoflagellates (NF) and ciliates. Protistan grazing impact on both heterotrophic bacteria and PCya was mainly due to NF, which contributed up to 96% of the microbial grazing. There was a clear selection of FLA by protists. PCya represented the main carbon source for both flagellates and ciliates in the mixolimnion, accounting for an average of 83% of the total carbon obtained from the ingestion of picoplanktonic organisms. Protists were the main consumers of particulate primary production (46-74% depending on season). Significant seasonal variation of grazing rates (0.011-0.041 h(-1)) was found, chiefly following variation of PCya production and biomass. Assuming a growth efficiency of 0.4, total protozoan production varied seasonally (189-313 g C m(-2) day(-1)) and was roughly half of particulate phytoplankton production. This study provides evidence that NF and PCya were tightly coupled in Lake Tanganyika and that herbivory by protists may be one of the reasons why this great lake has high productivity. Our results bring support to the idea that microbial herbivory is a major process in oligotrophic freshwater systems.

Prolonging the lifetime and activity of silica immobilised Cyanidium caldarium.

Over the past few years the idea of living photosynthetic materials has advanced from concept to reality. This work outlines the improvements made in the immobilisation of the thermotolerant acidophile Cyanidium caldarium (Tilden) Geitler SAG 16.91 within porous and transparent silica gels with the view to targeting photochemical materials that can be used to mitigate rising CO(2) emissions. Our results suggest that the immobilised cells are autofluorescent for at least 75 days post encapsulation and can maintain a steady oxygen production rate over a similar timeframe corroborating the viability and physiological activity of silica immobilised C. caldarium.

Vertical distribution of ammonia-oxidizing crenarchaeota and methanogens in the epipelagic waters of Lake Kivu (Rwanda-Democratic Republic of the Congo).

Four stratified basins in Lake Kivu (Rwanda-Democratic Republic of the Congo) were sampled in March 2007 to investigate the abundance, distribution, and potential biogeochemical role of planktonic archaea. We used fluorescence in situ hybridization with catalyzed-reported deposition microscopic counts (CARD-FISH), denaturing gradient gel electrophoresis (DGGE) fingerprinting, and quantitative PCR (qPCR) of signature genes for ammonia-oxidizing archaea (16S rRNA for marine Crenarchaeota group 1.1a [MCG1] and ammonia monooxygenase subunit A [amoA]). Abundance of archaea ranged from 1 to 4.5% of total DAPI (4',6-diamidino-2-phenylindole) counts with maximal concentrations at the oxic-anoxic transition zone (∼50-m depth). Phylogenetic analysis of the archaeal planktonic community revealed a higher level of richness of crenarchaeal 16S rRNA gene sequences (21 of the 28 operational taxonomic units [OTUs] identified [75%]) over euryarchaeotal ones (7 OTUs). Sequences affiliated with the kingdom Euryarchaeota were mainly recovered from the anoxic water compartment and mostly grouped into methanogenic lineages (Methanosarcinales and Methanocellales). In turn, crenarchaeal phylotypes were recovered throughout the sampled epipelagic waters (0- to 100-m depth), with clear phylogenetic segregation along the transition from oxic to anoxic water masses. Thus, whereas in the anoxic hypolimnion crenarchaeotal OTUs were mainly assigned to the miscellaneous crenarchaeotic group, the OTUs from the oxic-anoxic transition and above belonged to Crenarchaeota groups 1.1a and 1.1b, two lineages containing most of the ammonia-oxidizing representatives known so far. The concomitant vertical distribution of both nitrite and nitrate maxima and the copy numbers of both MCG1 16S rRNA and amoA genes suggest the potential implication of Crenarchaeota in nitrification processes occurring in the epilimnetic waters of the lake.

Strong effects of amoebae grazing on the biomass and genetic structure of a Microcystis bloom (Cyanobacteria).

Despite its importance for bloom toxicity, the factors determining the population structure of cyanobacterial blooms are poorly understood. Here, we report the results of a two-year field survey of the population dynamics of Microcystis blooms in a small hypertrophic urban pond. Microscopic enumeration of Microcystis and its predators and parasites was combined with pigment and microcystin analysis and denaturing gradient gel electrophoresis of the ITS rDNA region to assess population dynamics and structure. Two main Microcystis morpho- and ITS types were revealed, corresponding to M. aeruginosa and M. viridis. In both years, high population densities of naked amoebae grazing on Microcystis coincided with rapid decreases in Microcystis biomass. In one year, there was a shift from heavily infested M. aeruginosa to the less-infested M. viridis, allowing the bloom to rapidly recover. The preference of amoebae for M. aeruginosa was confirmed by grazing experiments, in which several amoeba strains were capable of grazing down a strain of M. aeruginosa, but not of M. viridis. Zooplankton and chytrid parasites appeared to be of minor importance for these strong and fast reductions in Microcystis biomass. These findings demonstrate a strong impact of small protozoan grazers on the biomass and genetic structure of Microcystis blooms.

Hybrid photosynthetic materials derived from microalgae Cyanidium caldarium encapsulated within silica gel.

Cyanidium caldarium (Tilden) Geitler SAG 16.91 has been encapsulated within a porous silica host structure to target novel photosynthetic hybrid materials suitable for use in solar cells or CO(2) fixation. C. caldarium cells are both thermophilic and acidophilic; on account of these tolerances the hybrid materials could be employed in more extreme heat conditions. TEM highlights that the external cell membrane can remain intact after encapsulation. The images reveal an alignment of silica gel around the external membrane of the cell, providing evidence that the cell wall acts as both a nucleation and polymerisation site for silica species and that the silica scaffold formed by the aggregation of colloidal particles, generates a porosity that can facilitate the transport of nutrients towards the cell. Epifluorescence microscopy and UV-visible spectroscopy have revealed the preservation of photosynthetic apparatus post-immobilisation. Productivity studies showed how the presence of silica nanoparticles within the matrix can adversely interact with the exterior cellular structures preventing the production of oxygen through photosynthesis.

Fate of heterotrophic bacteria in Lake Tanganyika (East Africa).

Bacterial mortality was studied using two complementary methods between 2002 and 2004 in the two main basins (north and south) of Lake Tanganyika. The disappearance of radioactivity from the DNA of natural assemblages of bacteria previously labeled with tritiated thymidine was used to estimate the mortality due to grazing by predators (72%) and due to the cell lysis (28%). Measurements of ingestion rate of bacteria by protozoa using fluorescent micro-particles yielded protozoan grazing rates similar to those provided by the thymidine method, and showed that heterotrophic nano-flagellates were responsible for most of the grazing pressure on the bacterial community of the pelagic zone (92-99%). Bacterial cell lysis was the second process involved in bacterial mortality, ranking before ciliate grazing. Overall, bacterial mortality was balanced with bacterial production. With regard to the assessment of the trophic role of bacteria, it was estimated that c. 5-8% of the organic carbon taken up by bacteria was converted into protozoan biomass and was thus available for metazoans.

Bacterial community composition in Lake Tanganyika: vertical and horizontal heterogeneity.

Vertical and latitudinal differences in bacterial community composition (BCC) in Lake Tanganyika were studied during the dry season of 2002 by means of denaturing gradient gel electrophoresis analysis of PCR-amplified 16S RNA fragments. Dominant bands were sequenced and identified as members of the Cyanobacteria, Actinobacteria, Nitrospirae, green nonsulfur bacteria, and Firmicutes divisions and the Gamma- and Deltaproteobacteria subdivisions. The BCC in the lake displayed both vertical and latitudinal variation. Vertical changes in BCC were related to the thermal water column stratification, which influences oxygen and nutrient concentrations. Latitudinal variation was related to upwelling of deep water and increased primary production in the south of the lake. The number of bands per sample increased with bacterial production in the epilimnion of the lake, suggesting a positive diversity-productivity relationship.